Article | Published:

Unique treatment potential of cannabidiol for the prevention of relapse to drug use: preclinical proof of principle

Neuropsychopharmacologyvolume 43pages20362045 (2018) | Download Citation


Cannabidiol (CBD), the major non-psychoactive constituent of Cannabis sativa, has received attention for therapeutic potential in treating neurologic and psychiatric disorders. Recently, CBD has also been explored for potential in treating drug addiction. Substance use disorders are chronically relapsing conditions and relapse risk persists for multiple reasons including craving induced by drug contexts, susceptibility to stress, elevated anxiety, and impaired impulse control. Here, we evaluated the “anti-relapse” potential of a transdermal CBD preparation in animal models of drug seeking, anxiety and impulsivity. Rats with alcohol or cocaine self-administration histories received transdermal CBD at 24 h intervals for 7 days and were tested for context and stress-induced reinstatement, as well as experimental anxiety on the elevated plus maze. Effects on impulsive behavior were established using a delay-discounting task following recovery from a 7-day dependence-inducing alcohol intoxication regimen. CBD attenuated context-induced and stress-induced drug seeking without tolerance, sedative effects, or interference with normal motivated behavior. Following treatment termination, reinstatement remained attenuated up to ≈5 months although plasma and brain CBD levels remained detectable only for 3 days. CBD also reduced experimental anxiety and prevented the development of high impulsivity in rats with an alcohol dependence history. The results provide proof of principle supporting potential of CBD in relapse prevention along two dimensions: beneficial actions across several vulnerability states and long-lasting effects with only brief treatment. The findings also inform the ongoing medical marijuana debate concerning medical benefits of non-psychoactive cannabinoids and their promise for development and use as therapeutics.

Access optionsAccess options

Rent or Buy article

Get time limited or full article access on ReadCube.


All prices are NET prices.


  1. 1.

    Wilkinson ST, Yarnell S, Radhakrishnan R, Ball SA, D’Souza DC. Marijuana legalization: impact on physicians and public health. Annu Rev Med. 2016;67:453–66.

  2. 2.

    Hurd YL. Cannabidiol: swinging the marijuana pendulum from ‘Weed’ to medication to treat the opioid epidemic. Trends Neurosci. 2017;40:124–7.

  3. 3.

    Scuderi C, De Filippis D, Iuvone T, Blasio A, Steardo A, Esposito G. Cannabidiol in medicine: a review of its therapeutic potential in CNS disorders. Phytother Res. 2009;23:597–602.

  4. 4.

    O’Connell BK, Gloss D, Devinsky O. Cannabinoids in treatment-resistant epilepsy: a review. Epilepsy Behav. 2017;70(Pt B):341–8.

  5. 5.

    Russo M, Naro A, Leo A, Sessa E, D’Aleo G, Bramanti P, et al. Evaluating sativex(R) in neuropathic pain management: a clinical and neurophysiological assessment in multiple sclerosis. Pain Med. 2016;17:1145–54.

  6. 6.

    Consroe P, Carlini EA, Zwicker AP, Avelinolacerda L. Interaction of cannabidiol and alcohol in humans. Psychopharmacology. 1979;66:45–50.

  7. 7.

    Morgan CJ, Das RK, Joye A, Curran HV, Kamboj SK. Cannabidiol reduces cigarette consumption in tobacco smokers: preliminary findings. Addict Behav. 2013;38:2433–6.

  8. 8.

    Katsidoni V, Anagnostou I, Panagis G. Cannabidiol inhibits the reward-facilitating effect of morphine: involvement of 5-HT1A receptors in the dorsal raphe nucleus. Addict Biol. 2013;18:286–96.

  9. 9.

    Mahmud A, Gallant S, Sedki F, D’Cunha T, Shalev U. Effects of an acute cannabidiol treatment on cocaine self-administration and cue-induced cocaine seeking in male rats. J Psychopharmacol. 2016;31:96–104.

  10. 10.

    Parker LA, Burton P, Sorge RE, Yakiwchuk C, Mechoulam R. Effect of low doses of delta9-tetrahydrocannabinol and cannabidiol on the extinction of cocaine-induced and amphetamine-induced conditioned place preference learning in rats. Psychopharmacology. 2004;175:360–6.

  11. 11.

    Ren YH, Whittard J, Higuera-Matas A, Morris CV, Hurd YL. Cannabidiol, a nonpsychotropic component of cannabis, inhibits cue-induced heroin seeking and normalizes discrete mesolimbic neuronal disturbances. J Neurosci. 2009;29:14764–9.

  12. 12.

    Prud’homme M, Cata R, Jutras-Aswad D. Cannabidiol as an intervention for addictive behaviors: a systematic review of the evidence. Subst Abus. 2015;9:33–8.

  13. 13.

    American Psychiatric Association. Diagnostic and statistical manual of mental disorders. 5th ed. Arlington, VA: American Psychiatric Publishing; 2013.

  14. 14.

    Leshner AI. Addiction is a brain disease, and it matters. Science. 1997;278:45–7.

  15. 15.

    Campos AC, Ferreira FR, Guimaraes FS. Cannabidiol blocks long-lasting behavioral consequences of predator threat stress: possible involvement of 5HT1A receptors. J Psychiatr Res. 2012;46:1501–10.

  16. 16.

    Guimaraes FS, Chiaretti TM, Graeff FG, Zuardi AW. Antianxiety effect of cannabidiol in the elevated plus-maze. Psychopharmacology. 1990;100:558–9.

  17. 17.

    Campos AC, Ortega Z, Palazuelos J, Fogaca MV, Aguiar DC, Diaz-Alonso J, et al. The anxiolytic effect of cannabidiol on chronically stressed mice depends on hippocampal neurogenesis: involvement of the endocannabinoid system. Int J Neuropsychopharmacol. 2013;16:1407–19.

  18. 18.

    Marinho AL, Vila-Verde C, Fogaca MV, Guimaraes FS. Effects of intra-infralimbic prefrontal cortex injections of cannabidiol in the modulation of emotional behaviors in rats: contribution of 5HT(1)A receptors and stressful experiences. Behav Brain Res. 2015;286:49–56.

  19. 19.

    Zanelati TV, Biojone C, Moreira FA, Guimaraes FS, Joca SRL. Antidepressant-like effects of cannabidiol in mice: possible involvement of 5-HT(1A) receptors. Br J Pharmacol. 2010;159:122–8.

  20. 20.

    Casarotto PC, Gomes FV, Resstel LB, Guimaraes FS. Cannabidiol inhibitory effect on marble-burying behaviour: involvement of CB1 receptors. Behav Pharmacol. 2010;21:353–8.

  21. 21.

    Gomes FV, Reis DG, Alves FHF, Correa FMA, Guimaraes FS, Resstel LBM. Cannabidiol injected into the bed nucleus of the stria terminalis reduces the expression of contextual fear conditioning via 5-HT1A receptors. J Psychopharmacol. 2012;26:104–13.

  22. 22.

    Guimaraes VM, Zuardi AW, Del Bel EA, Guimaraes FS. Cannabidiol increases Fos expression in the nucleus accumbens but not in the dorsal striatum. Life Sci. 2004;75:633–8.

  23. 23.

    Lemos JI, Resstel LB, Guimaraes FS. Involvement of the prelimbic prefrontal cortex on cannabidiol-induced attenuation of contextual conditioned fear in rats. Behav Brain Res. 2010;207:105–11.

  24. 24.

    Koob GF, Volkow ND. Neurocircuitry of addiction. Neuropsychopharmacology. 2010;35:217–38.

  25. 25.

    Iuvone T, Esposito G, De Filippis D, Scuderi C, Steardo L. Cannabidiol: a promising drug for neurodegenerative disorders? Cns Neurosci & Ther. 2009;15:65–75.

  26. 26.

    Liput DJ, Hammell DC, Stinchcomb AL, Nixon K. Transdermal delivery of cannabidiol attenuates binge alcohol-induced neurodegeneration in a rodent model of an alcohol use disorder. Pharmacol Biochem Behav. 2013;111:120–7.

  27. 27.

    Crews FT, Boettiger CA. Impulsivity, frontal lobes and risk for addiction. Pharmacol Biochem Behav. 2009;93:237–47.

  28. 28.

    Agurell S, Carlsson S, Lindgren JE, Ohlsson A, Gillespie H, Hollister L. Interactions of delta 1-tetrahydrocannabinol with cannabinol and cannabidiol following oral administration in man. Assay of cannabinol and cannabidiol by mass fragmentography. Experientia. 1981;37:1090–2.

  29. 29.

    Merrick J, Lane B, Sebree T, Yaksh T, O’Neill C, Banks SL. Identification of psychoactive degradants of cannabidiol in simulated gastric and physiological fluid. Cannabis Cannabinoid Res. 2016;1:102–13.

  30. 30.

    Paudel KS, Hammell DC, Agu RU, Valiveti S, Stinchcomb AL. Cannabidiol bioavailability after nasal and transdermal application: effect of permeation enhancers. Drug Dev Ind Pharm. 2010;36:1088–97.

  31. 31.

    Kufahl PR, Martin-Fardon R, Weiss F. Enhanced sensitivity to attenuation of conditioned reinstatement by the mGluR(2/3) agonist LY379268 and increased functional activity of mGluR(2/3) in rats with a history of ethanol dependence. Neuropsychopharmacology. 2011;36:2762–73.

  32. 32.

    Martin-Fardon R, Baptista MA, Dayas CV, Weiss F. Dissociation of the effects of MTEP [3-[(2-methyl-1,3-thiazol-4-yl)ethynyl]piperidine] on conditioned reinstatement and reinforcement: comparison between cocaine and a conventional reinforcer. J Pharmacol Exp Ther. 2009;329:1084–90.

  33. 33.

    Zhao Y, Weiss F, Zorrilla EP. Remission and resurgence of anxiety-like behavior across protracted withdrawal stages in ethanol-dependent rats. Alcohol Clin Exp Res. 2007;31:1505–15.

  34. 34.

    Pattij T, Janssen MC, Schepers I, Gonzalez-Cuevas G, de Vries TJ, Schoffelmeer AN. Effects of the cannabinoid CB1 receptor antagonist rimonabant on distinct measures of impulsive behavior in rats. Psychopharmacology. 2007;193:85–96.

  35. 35.

    Baptista MA, Martin-Fardon R, Weiss F. Preferential effects of the metabotropic glutamate 2/3 receptor agonist LY379268 on conditioned reinstatement versus primary reinforcement: comparison between cocaine and a potent conventional reinforcer. J Neurosci. 2004;24:4723–7.

  36. 36.

    Ciccocioppo R, Martin-Fardon R, Weiss F. Stimuli associated with a single cocaine experience elicit long-lasting cocaine-seeking. Nat Neurosci. 2004;7:495–6.

  37. 37.

    Ciccocioppo R, Sanna PP, Weiss F. Cocaine-predictive stimulus induces drug-seeking behavior and neural activation in limbic brain regions after multiple months of abstinence: reversal by D(1) antagonists. Proc Natl Acad Sci USA. 2001b;98:1976–81.

  38. 38.

    Martin-Fardon R, Weiss F. Perseveration of craving: effects of stimuli conditioned to drugs of abuse versus conventional reinforcers differing in demand. Addict Biol. 2017;22:923–32.

  39. 39.

    Weiss F, Maldonado-Vlaar CS, Parsons LH, Kerr TM, Smith DL, Ben-Shahar O. Control of cocaine-seeking behavior by drug-associated stimuli in rats: effects on recovery of extinguished operant-responding and extracellular dopamine levels in amygdala and nucleus accumbens. Proc Natl Acad Sci USA. 2000;97:4321–6.

  40. 40.

    Weiss F, Martin-Fardon R, Ciccocioppo R, Kerr TM, Smith DL, Ben-Shahar O. Enduring resistance to extinction of cocaine-seeking behavior induced by drug-related cues. Neuropsychopharmacology. 2001;25:361–72.

  41. 41.

    Ciccocioppo R, Angeletti S, Weiss F. Long-lasting resistance to extinction of response reinstatement induced by ethanol-related stimuli: role of genetic ethanol preference. Alcohol Clin Exp Res. 2001;25:1414–9.

  42. 42.

    Katner SN, Magalong JG, Weiss F. Reinstatement of alcohol-seeking behavior by drug-associated discriminative stimuli after prolonged extinction in the rat. Neuropsychopharmacology. 1999;20:471–9.

  43. 43.

    Katner SN, Weiss F. Ethanol-associated olfactory stimuli reinstate ethanol-seeking behavior after extinction and modify extracellular dopamine levels in the nucleus accumbens. Alcohol Clin Exp Res. 1999;23:1751–60.

  44. 44.

    Liu X, Weiss F. Reversal of ethanol-seeking behavior by D1 and D2 antagonists in an animal model of relapse: differences in antagonist potency in previously ethanol-dependent versus nondependent rats. J Pharmacol Exp Ther. 2002;300:882–9.

  45. 45.

    Sidhpura N, Weiss F, Martin-Fardon R. Effects of the mGlu2/3 agonist LY379268 and the mGlu5 antagonist MTEP on ethanol seeking and reinforcement are differentially altered in rats with a history of ethanol dependence. Biol Psychiatry. 2010;67:804–11.

  46. 46.

    Crombag HS, Bossert JM, Koya E, Shaham Y. Context-induced relapse to drug seeking: a review. Philos Trans R Soc Lond B Biol Sci. 2008;363:3233–43.

  47. 47.

    Aujla H, Cannarsa R, Romualdi P, Ciccocioppo R, Martin-Fardon R, Weiss F. Modification of anxiety-like behaviors by nociceptin/orphanin FQ (N/OFQ) and time-dependent changes in N/OFQ-NOP gene expression following ethanol withdrawal. Addict Biol. 2013;18:467–79.

  48. 48.

    Braconi S, Sidhpura N, Aujla H, Martin-Fardon R, Weiss F, Ciccocioppo R. Revisiting intragastric ethanol intubation as a dependence induction method for studies of ethanol reward and motivation in rats. Alcohol Clin Exp Res. 2010;34:538–44.

  49. 49.

    Folch-Pi J, Lees M, Stanley GHS. A simple method for the isolation and purification of total lipids from animal tissue. J Biol Chem. 1957;226:497–509.

  50. 50.

    Martin-Fardon R, Weiss F. Modeling relapse in animals. Curr Top Behav Neurosci. 2013;13:403–32.

  51. 51.

    Chen YW, Fiscella KA, Bacharach SZ, Tanda G, Shaham Y, Calu DJ. Effect of yohimbine on reinstatement of operant responding in rats is dependent on cue contingency but not food reward history. Addict Biol. 2015;20:690–700.

  52. 52.

    Dick DM, Smith G, Olausson P, Mitchell SH, Leeman RF, O’Malley SS, et al. Understanding the construct of impulsivity and its relationship to alcohol use disorders. Addict Biol. 2010;15:217–26.

  53. 53.

    Verdejo-Garcia A, Lawrence AJ, Clark L. Impulsivity as a vulnerability marker for substance-use disorders: Review of findings from high-risk research, problem gamblers and genetic association studies. Neurosci Biobehav Rev. 2008;32:777–810.

  54. 54.

    Stevens L, Goudriaan AE, Verdejo-Garcia A, Dom G, Roeyers H, Vanderplasschen W. Impulsive choice predicts short-term relapse in substance-dependent individuals attending an in-patient detoxification programme. Psychol Med. 2015;45:2083–93.

  55. 55.

    Oberlin BG, Grahame NJ. High-alcohol preferring mice are more impulsive than low-alcohol preferring mice as measured in the delay discounting task. Alcohol Clin Exp Res. 2009;33:1294–303.

  56. 56.

    Poulos CX, Parker JL, Le DA. Increased impulsivity after injected alcohol predicts later alcohol consumption in rats: evidence for “loss-of-control drinking” and marked individual differences. Behav Neurosci. 1998;112:1247–57.

  57. 57.

    Wilhelm CJ, Mitchell SH. Rats bred for high alcohol drinking are more sensitive to delayed and probabilistic outcomes. Genes Brain Behav. 2008;7:705–13.

  58. 58.

    Martin-Fardon R, Weiss F. Modeling relapse in animals. Curr Top Behav Neurosci. 2013;13:403–432.

  59. 59.

    Mantsch JR, Baker DA, Funk D, Le AD, Shaham Y. Stress-induced reinstatement of drug seeking: 20 years of progress. Neuropsychopharmacology. 2016;41:335–56.

  60. 60.

    Epstein DH, Preston KL, Stewart J, Shaham Y. Toward a model of drug relapse: an assessment of the validity of the reinstatement procedure. Psychopharmacology. 2006;189:1–16.

  61. 61.

    de Carvalho CR, Takahashi RN. Cannabidiol disrupts the reconsolidation of contextual drug-associated memories in Wistar rats. Addict Biol. 2017;22:742–51.

  62. 62.

    Crippa JAS, Derenusson GN, Ferrari TB, Wichert-Ana L, Duran FLS, Martin-Santos R, et al. Neural basis of anxiolytic effects of cannabidiol (CBD) in generalized social anxiety disorder: a preliminary report. J Psychopharmacol. 2011;25:121–30.

  63. 63.

    Bergamaschi MM, Queiroz RHC, Chagas MHN, de Oliveira DCG, De Martinis BS, Kapczinski F, et al. Cannabidiol reduces the anxiety induced by simulated public speaking in treatment-naive social phobia patients. Neuropsychopharmacology. 2011;36:1219–26.

  64. 64.

    Stern CA, Gazarini L, Takahashi RN, Guimaraes FS, Bertoglio LJ. On disruption of fear memory by reconsolidation blockade: evidence from cannabidiol treatment. Neuropsychopharmacology. 2012;37:2132–42.

  65. 65.

    Arndt DL, de Wit H. Cannabidiol does not dampen responses to emotional stimuli in healthy adults. Cannabis Cannabinoid Res. 2017;2:105–13.

  66. 66.

    ElBatsh MM, Assareh N, Marsden CA, Kendall DA. Anxiogenic-like effects of chronic cannabidiol administration in rats. Psychopharmacology. 2012;221:239–47.

  67. 67.

    Campos AC, Moreira FA, Gomes FV, Del Bel EA, Guimaraes FS. Multiple mechanisms involved in the large-spectrum therapeutic potential of cannabidiol in psychiatric disorders. Philos Trans R Soc Lond B Biol Sci. 2012;367:3364–78.

  68. 68.

    Parsons LH, Hurd YL. Endocannabinoid signalling in reward and addiction. Nat Rev Neurosci. 2015;16:579–94.

  69. 69.

    Ashton CH, Moore PB. Endocannabinoid system dysfunction in mood and related disorders. Acta Psychiatr Scand. 2011;124:250–61.

  70. 70.

    Moreira FA, Wotjak CT. Cannabinoids and anxiety. Curr Top Behav Neurosci. 2010;2:429–50.

  71. 71.

    Riebe CJ, Wotjak CT. Endocannabinoids and stress. Stress. 2011;14:384–97.

  72. 72.

    Kathmann M, Flau K, Redmer A, Trankle C, Schlicker E. Cannabidiol is an allosteric modulator at mu- and delta-opioid receptors. Naunyn Schmiede Arch Pharmacol. 2006;372:354–61.

  73. 73.

    Gururajan A, Taylor DA, Malone DT. Cannabidiol and clozapine reverse MK-801-induced deficits in social interaction and hyperactivity in Sprague-Dawley rats. J Psychopharmacol. 2012;26:1317–32.

  74. 74.

    Russo EB, Burnett A, Hall B, Parker KK. Agonistic properties of cannabidiol at 5-HT1a receptors. Neurochem Res. 2005;30:1037–43.

  75. 75.

    Campos AC, Guimaraes FS. Involvement of 5HT1A receptors in the anxiolytic-like effects of cannabidiol injected into the dorsolateral periaqueductal gray of rats. Psychopharmacology. 2008;199:223–30.

  76. 76.

    Fogaca MV, Reis FM, Campos AC, Guimaraes FS. Effects of intra-prelimbic prefrontal cortex injection of cannabidiol on anxiety-like behavior: involvement of 5HT1A receptors and previous stressful experience. Eur Neuropsychopharmacol. 2014;24:410–9.

  77. 77.

    Gomes FV, Resstel LBM, Guimaraes FS. The anxiolytic-like effects of cannabidiol injected into the bed nucleus of the stria terminalis are mediated by 5-HT1A receptors. Psychopharmacology. 2011;213:465–73.

  78. 78.

    Resstel LB, Tavares RF, Lisboa SF, Joca SR, Correa FM, Guimaraes FS. 5-HT1A receptors are involved in the cannabidiol-induced attenuation of behavioural and cardiovascular responses to acute restraint stress in rats. Br J Pharmacol. 2009;156:181–8.

  79. 79.

    Mandyam CD, Koob GF. The addicted brain craves new neurons: putative role for adult-born progenitors in promoting recovery. Trends Neurosci. 2012;35:250–60.

  80. 80.

    Jiang W, Zhang Y, Xiao L, Van Cleemput J, Ji SP, Bai G, et al. Cannabinoids promote embryonic and adult hippocampus neurogenesis and produce anxiolytic- and antidepressant-like effects. J Clin Invest. 2005;115:3104–16.

  81. 81.

    Wolf SA, Bick-Sander A, Fabel K, Leal-Galicia P, Tauber S, Ramirez-Rodriguez G, et al. Cannabinoid receptor CB1 mediates baseline and activity-induced survival of new neurons in adult hippocampal neurogenesis. Cell Commun Signal. 2010;8:12.

  82. 82.

    Amlung M, Vedelago L, Acker J, Balodis I, MacKillop J. Steep delay discounting and addictive behavior: a meta-analysis of continuous associations. Addiction. 2017;112:51–62.

  83. 83.

    MacKillop J, Kahler CW. Delayed reward discounting predicts treatment response for heavy drinkers receiving smoking cessation treatment. Drug Alcohol Depend. 2009;104:197–203.

  84. 84.

    Crews FT, Boettiger CA. Impulsivity, frontal lobes and risk for addiction. Pharmacol Biochem Behav. 2009; 93:237–247.

  85. 85.

    Hampson AJ, Grimaldi M, Axelrod J, Wink D. Cannabidiol and (-)delta(9)-tetrahydrocannabinol are neuroprotective antioxidants. Proc Natl Acad Sci USA. 1998;95:8268–73.

  86. 86.

    Nixon K, Crews FT. Binge ethanol exposure decreases neurogenesis in adult rat hippocampus. J Neurochem. 2002;83:1087–93.

  87. 87.

    Esposito G, Scuderi C, Valenza M, Togna GI, Latina V, De Filippis D, et al. Cannabidiol reduces Abeta-induced neuroinflammation and promotes hippocampal neurogenesis through PPARgamma involvement. PLoS ONE. 2011;6:e28668.

  88. 88.

    Fagherazzi EV, Garcia VA, Maurmann N, Bervanger T, Halmenschlager LH, Busato SB, et al. Memory-rescuing effects of cannabidiol in an animal model of cognitive impairment relevant to neurodegenerative disorders. Psychopharmacology. 2012;219:1133–40.

Download references


This is publication number 29366 from the Scripps Research Institute. The CBD gel preparation was generously provided by Zynerba Pharmaceuticals, Inc, (Devon, PA) and AllTranz, Inc. (Lexington, KY). We thank the students Christine Chan, William Choy, Yunchen Ding, Li Huang, Tao Moua, Mark Sanderson-Cimino, and Angela Wong for assistance.

Author contributions

FW and ALS conceived the study. GGC, FW, and RMF designed the experiments. GGC and FW analyzed the data and wrote the manuscript. GGC and RMF conducted the behavioral tests. SLB developed and prepared the transdermal CBD gel formulation. TMK, DGS, and LHP performed tissue extraction and LC-MS assays of CBD brain levels. DCH performed the LC-MS assays of CBD plasma levels.

Author information

Author notes

    • Gustavo Gonzalez-Cuevas

    Present address: Department of Psychology, European University of Madrid, School of Biomedical Sciences, Madrid, 28670, Spain


  1. Department of Neuroscience, The Scripps Research Institute, 10550 North Torrey Pines Road (SP30-2120), La Jolla, CA, 92037, USA

    • Gustavo Gonzalez-Cuevas
    • , Remi Martin-Fardon
    • , Tony M. Kerr
    • , David G. Stouffer
    • , Loren H. Parsons
    •  & Friedbert Weiss
  2. University of Maryland, School of Pharmacy, Baltimore, MD, 21201, USA

    • Dana C. Hammell
    •  & Audra L. Stinchcomb
  3. Zynerba Pharmaceuticals, Devon, PA, 19333, USA

    • Stan L. Banks


  1. Search for Gustavo Gonzalez-Cuevas in:

  2. Search for Remi Martin-Fardon in:

  3. Search for Tony M. Kerr in:

  4. Search for David G. Stouffer in:

  5. Search for Loren H. Parsons in:

  6. Search for Dana C. Hammell in:

  7. Search for Stan L. Banks in:

  8. Search for Audra L. Stinchcomb in:

  9. Search for Friedbert Weiss in:

Conflict of interest

The authors declare that they have no conflict of interest.

Corresponding author

Correspondence to Friedbert Weiss.

Electronic supplementary material

About this article

Publication history